8.3 Interface Classes and Objects
8.3.3 Method Inheritance
Uno nunca desea hacer estragos en las poblaciones más allá de lo necesario para satisfacer las necesidades de un estu- dio científico. Afortunadamente, coleccionar obreras de co- lonias grandes es como cortar o podar un arbusto o extraer unas cuantas células de la piel: por lo general uno deja la colonia (reina y obreras) poco alterada detrás de sí. Escoja un periodo de colección que maximice la actividad de las hormigas, típicamente uno donde las temperaturas sean altas y las lluvias abundantes. Las comunidades de desierto y trópico frecuentemente son más activas durante la esta- ción húmeda. En las zonas templadas el comienzo de verano es un buen tiempo. En los climas mediterráneos, donde más llueve en invierno y es más cálido en verano, puede ser me- jor coleccionar en primavera. Evite coleccionar cuando hay agua presente en abundancia o la vegetación está mojada. Esto frena la actividad de hormigas, especialmente de las pequeñas.
Sea cuidadoso con el manejo de cebos, trampas de caída y cualquier cosa que pueda estar en contacto con las hormi- gas. Evite perfumes, colonias y repelentes.
Si un objetivo es hacer monitoreo de densidades de colo- nias, no se deben contar más de una entrada en nidos con múltiples entradas. Para evitar errores en este sobreconteo, se pueden ignorar nidos de la misma especie a una distancia dada (como un metro). También se pueden hacer “experi- mentos de trasplante”: si una hormiga puesta en una entrada de un nido es atacada vigorosamente, es probable que se trate de una hormiga de otra colonia.
Si va a usar cebos en sus protocolos, use pequeños, que sean difíciles de monoplozar por colonias grandes, agresi- vas. Por ejemplo, pedacitos de galletas (hechas de harina, azúcar, huevos y nueces) atraen la mayor variedad de hormi- gas, en comparación de mantequilla de maní, atún y agua azucarada. Aún las hormigas cortadoras de hojas (Attini) y algunas Dacetini cazadoras de colémbolos (Strumigenys,
Pyramica) se detienen para llevarse migajas de esas galle-
tas. En el oscuro suelo de un bosque tropical un investiga- dor puede ver la migaja moviéndose sobre el suelo antes de ver la hormiga.
Si es posible, expanda su esfuerzo de colección a lo largo del día. Una forma de hacer esto es usar técnicas de trampas pasivas como trampas de caída (pitfall).
La distribución en parche es un problema cuando se está ensamblando un programa de monitoreo. Las densidades de hormigas pueden variar mucho de un metro cuadrado a otro o de un valle a otro, por lo cual es importante determinar cuadrados de muestreo al azar y cuantos sean posibles para cubrir esta variabilidad.
En resumen, la ecología de hormigas, pequeñas, sésiles, co- loniales, organismos quimiosensoriales, difiere básicamente de la de los vertebrados y de ahí de nuestro propio mundo. El impacto de las hormigas en los ecosistemas es profundo y su presencia es ubicua. Por estas razones es una buena decisión incluir hormigas en las listas de taxa para progra- mas de monitoreo. Una cuidadosa consideración de la his- toria de vida de las hormigas, durante la fase de diseño, puede maximizar los dividendos de esta inversión.
Andersen, A. N. 1992. The rainforest ant fauna of the northern Kimberley region of Western Australia (Hymenoptera: Formicidae). Journal of the Australian
Entomological Society 31:187-192.
Andersen, A. N. 2000. A Global Ecology of Rainforest Ants: Functional groups in relation to environmental stress and disturbance, pp.25-34 in D. Agosti, J. D. Majer, L. E. Alonso y T. R. Schultz, eds., Ants: Standard Methods
for Measuring and Monitoring Biodiversity.
Smithsonian Institution Press.
Andersen, A. N. y A. D. Patel. 1994. Meat ants as dominant members of Australian ant communities: An experimen- tal test of their influence on the foraging success and forager abundance of other species. Oecologia 98:15- 24.
Bernstein, R. A. 1979. Schedules of foraging activity in species of ants. Journal of Animal Ecology 48:921-930. Beshers, S. y J. Traniello. 1994. The adaptiveness of worker
demography in the attine ant Trachymyrmex
Davidson, D. W. 1980. Some consequences of diffuse competition in a desert ant community. American
Naturalist 116:92-105.
Davidson, D. W. 1997. The role of resource imbalances in the evolutionary ecology of tropical arboreal ants.
Biological Journal of the Linnean Society 61:153-
181.
Davidson, D. W. 1998. Resource discovery versus resource domination in ants: A functional mechanism for breaking the tradeoff. Ecological Entomology 23:484-490. Davidson, D. W., J. T. Longino y R. R. Snelling. 1988. Pruning
of host ant neighbors by ants: An experimental approach. Ecology 69:801-808.
Davidson, D. W., R. R. Snelling y J. T. Longino. 1989. Competition among ants for myrmecophytes and the significance of plant trichomes. Biotropica 21:64-73. Deslippe, R. J. y R. Savolainen. 1994. Role of food supply in
structuring a population of Formica ants. Journal of
Animal Ecology 63:756-764.
Elmes, G. W. 1991. Ant colonies and environmental disturbance, pp.15-32 in: P. S. y A. Meadows, eds.,
Environmental Impact of Burrowing Animals and Animal Burrows. Clarendon Press, Oxford.
Erickson, J. M. 1971. The displacement of native ant species by the introduced Argentine ant Iridomyrmex humilis Mayr. Psyche 78:257-266.
Feener, D. H. Jr. 1981. Competition between ant species: Outcome controlled by parasitic flies. Science 214:815- 817.
Feener, D. H. Jr., L. F. Jacobs y J. O. Schmidt. 1996. Specialized parasitoid attracted to a pheromone of ants. Animal
Behavior 51:61-66.
Fellers, J. H. 1989. Daily and seasonal activity in woodlands ants. Oecologia 78:69-76.
Fernández, F. 1998. ¿Por qué hay tantas hormigas en los ár- boles?. Innovación y Ciencia 7(2):42-51.
Fittkau, E. J. y H. Klinge. 1973. On biomass and trophic structure of the Central Amazonian rain forest ecosystem. Biotropica 5:2-14.
Fluker, S. S. y J. W. Beardsley. 1970. Sympatric associations of three ants: Iridomyrmex humilis, Pheidole
megacephala, and Anoplolepis longiceps in Hawaii. Annals of the Entomological Society of America
63:1290-1296.
Fowler, H. G., J. V. E. Bernardi y L. F. T. di Romagnano. 1990. Community structure and Solenopsis invicta in São Paulo, pp.199-207 in: R. K. Vander Meer, K. Jaffé y A. Cedeno, eds., Applied Myrmecology: A World
Perspective Westview Press, Boulder.
Franks, N. R. y W. H. Bossert. 1983. The influence of swarm raiding army ants on the patchiness and diversity of a tropical leaf-litter ant community, pp.151-163 in: S. L. Sutton, T. C. Whitmore y A. C. Chadwick, eds., Tropi-
cal Rain Forest: Ecology and Management Blackwell,
Oxford.
Goldstein, E. L. 1975. Island biogeography of ants. Evolution 29:750-762.
Black, R. W. 1987. The biology of leaf nesting ants in a tropi- cal wet forest. Biotropica 19:319-325.
Blackburn, T., P. Harvey y M. Pagel. 1990. Species number, population density and body size relationships in na- tural communites. Journal of Animal Ecology 59:335- 345.
Bond, W. y P. Slingsby. 1984. Collapse of ant-plant mutualism: The argentine ant (Iridomyrmex humilis) and myrmecochorous proteaceae. Ecology 65:1031-1037. Brian, M. V. y A. D. Brian. 1951. Insolation and ant populations
in the west of Scotland. Transactions of the Royal
Entomological Society of London 102:303-330.
Brown, B. V. 1993. Taxonomy and preliminary phylogeny of the parasitic genus Apocephalus, subgenus
Mesophora (Diptera: Phoridae). Systematic Entomology 18:191-230.
Brown, B. V. y D. H. Feener. 1991a. Behavior and host location cues of Apocephalus paraponerae (Diptera: Phoridae), a parasitoid of the giant tropical ant, Paraponera
clavata (Hymenoptera: Formicidae). Biotropica 23:182-
187.
Brown, B. V. y D. H. Feener. 1991b. Life history parameters and description of the larva of Apocephalus paraponerae (Diptera: Phoridae), a parasitoid of the giant tropical ant, Paraponera clavata (Hymenoptera: Formicidae). Journal of Natural History 25:221-231. Brown, W. L. Jr. 1973. A comparison of the Hylean and Con-
go West African rain forest ant faunas, pp.161-185 in: B. J. Meggers, E. S. Ayensu y W. D. Duckworth, eds.,
Tropical Forest Ecosystems in Africa and South America: A comparative review Smithsonian
Institution Press, Washington D.C.
Byrne, M. M. 1994. Ecology of twig-dwelling ants in a wet lowland tropical forest. Biotropica 26:61-72.
Camilo, G. R. y S. A. Philips. 1990. Evolution of ant communities in response to invasion by the fire ant Solenopsis in-
victa, pp.190-198 in: R. K. Vander Meer, K. Jaffé y A.
Cedeno, eds., Applied Myrmecology: A World
Perspective Westview Press, Boulder.
Clark, D. B., C. Guayasamín, O. Pazmiño, C. Donoso y Y. Páez de Villacís. 1982. The tramp ant Wasmannia
auropunctata: Autecology and effects on ant diversity
and distribution on Santa Cruz Island. Galápagos
Biotropica 14:196-207.
Cushman, J. H., J. H. Lawton y B. F. J. Manly. 1993. Latitudi- nal patterns in European ants assemblages: Variation in species richness and body size. Oecologia 95:30-37. Davidson, D. W. 1977a. Species diversity and community organization in desert seed-eating ants. Ecology 58:711- 724.
Davidson, D. W. 1977b. Foraging ecology and community organization in desert seed-eating ants. Ecology 58:725- 737.
Davidson, D. W. 1978. Size variability in the worker caste of a social insect (Veromessor pergandei Mayr) as a function of the competitive environment. American
Gotelli, N. J. 1993. Ant lion zones: Causes of high-density predator aggregations. Ecology 74:226-237.
Greenslade, P. J. M. 1972. Comparative ecology of four tropi- cal ant species. Insectes Sociaux 19:195-212.
Haines, B. L. 1978. Element and energy flows through colonies of the leaf-cutting ant, Atta columbica, in Panamá.
Biotropica 10:270-277.
Haines, B. L. 1983. Leaf cutting-ants bleed mineral elements out of rainforest in southern Venezuela. Tropical
Ecology 24:85-93.
Hamilton, D. H. 1964. The genetical evolution of social behavior I. Journal of Theoretical Biology 7:1-16. Hamilton, D. H. 1972. Altruism and related phenomena, mainly
in social insects. Annual Review of Ecology and
Systematics 3:193-232.
Heinze, J. y B. Hölldobler. 1994. Ants in the cold. Memorabilia
Zoologica 48:99-108.
Herbers, J. M. 1985. Seasonal structuring of a north temperate ant community. Insectes Sociaux 32:224-240.
Herbers, J. M. 1989. Community structure in north temperate ants: Temporal and spatial variation. Oecologia 81:201- 211.
Hölldobler, B. 1983. Territorial behavior in the green ant (Oecophylla smaragdina). Biotropica 15:241-250. Hölldobler, B.y C. J. Lumsden. 1980. Territorial strategies in
ants. Science 210:732-739.
Hölldobler, B. y E. O. Wilson. 1977. The number of queens: an important trait in ant evolution. Naturwissenschaften 64:8-15.
Hölldobler, B. y E. O. Wilson. 1990. The Ants Harvard University Press, Cambridge, 732 pp.
Hood, W. G. y W. R. Tschinkel. 1990. Dessication resistance in arboreal and terrestrial ants. Physiological
Entomology 15:23-35.
Huxley, C. R. y D. F. Cutler, eds. 1991. Ant-Plant Interactions Oxford University Press, Oxford.
Jeanne, R. L. 1979. A latitudinal gradient in rates of ant predation. Ecology 60:1211-1224.
Johnson, R. A. 1992. Soil texture as an influence on the distribution of the desert seed-harvest ants
Pogonomyrmex rugosus and Messor pergandei. Oecologia 89:118-124.
Kaspari, M. 1993a. Body size and microclimate use in Neotropical granivorous ants. Oecologia 96:500-507. Kaspari, M. 1993b. Removal of seeds from Neotropical
frugivore droppings. Oecologia 95:81-99.
Kaspari, M. 1996a. Litter ant patchiness at the 1 m2 scale:
Disturbance dynamics in three Neotropical forests.
Oecologia 107:265-273.
Kaspari, M. 1996b. Testing resource-based models of patchiness in four Neotropical litter ant assemblages.
Oikos 76:443-454.
Kaspari, M. 1996c. Worker size and seed size selection by harvester ants in a Neotropical forest. Oecologia 105:397-404.
Kaspari, M. y M. Byrne. 1995. Caste allocation in litter
Pheidole. Behavioral Ecology and Sociobiology
37:255-263.
Kaspari, M. y E. L. Vargo. 1995. Colony size as a buffer against seasonality: Bergmann’s rule in social insects.
American Naturalists 145:610-632.
Kaspari, M. y M. Weiser. 1999. The size-grain hypothesis and inbterspecific scaling in ants. Functional Ecology 13:530-538.
Kaspari, M., L. Alonso y S. O’Donnel. 2000a. Three energy variables predict ant abundance at a geographic scale.
Proceedings of the Royal Society B 267:485-490.
Kaspari, M., S. O’Donnel y J. Kercher. 2000b Energy, density, and constraints to species richness: Studies of ant assemblages along a productivity gradient. American
Naturalist 155:280-293.
Kaspari, M. y S. P. Yanoviak. 2000c. Bait use in tropical litter and canopy ants – evidence for differences in nutriton limitation. Biotropica 33(1):207-211.
Kaspari, M., S. O’Donnell y J. Kercher. 2000. Energy, density, and constraints to species richness: Studies of ant assemblages along a productivity gradient. American
Naturalist 155:280-293.
Kistner, D. H. 1982. The Social Insects’ Bestiary, pp.1-244 in: H. R. Hermann, ed., Social Insects Vol 3. Academic Press, New York.
Lawton, J. 1994. What do species do in ecosystems? Oikos 71:364-374.
Lesica, P. y P. Kannowski. 1998. Ants create hummocks and alter structure and vegetation of a mountain fen.
American Midland Naturalist 139:58-68.
Lévieux, J. 1976. Étude de la structure du nid de quelques espéces terricoles de fourmis tropicales. Annales de
l’Université d’Abidjan, Serie C: Sciences 12:23-33.
Lévieux, J. 1983. The soil fauna of tropical savannas IV The ants, pp.525-540 in: F. Bourliére, ed., Tropical
Savannas: Ecosystems of the World Elsvier,
Amsterdam.
Levings, S. 1983. Seasonal, annual and among-site variation in the ground ant community of a deciduous tropical forest: Some causes of patchy species distribution.
Ecological Monographs 53:435-455.
Levings, S. y J. F. A. Traniello. 1981. Territoriality, nest dispersion, and community structure in ants. Psyche 88:265-319.
Longino, J. T. 1991. Azteca ants in Cecropia trees: Taxonomy, colony structure, and behavior, pp.271-288 in: C. R. Huxley y F. D. Cutler, eds., Ant-Plant Interactions Oxford University Press, Oxford.
Lubin, Y. D. 1984. Changes in teh native ant fauna of the Galápagos Islands following invasion by the little red fire ant, Wasmannia auropunctata. Biological Journal
of the Linnean Society 21:229-242.
Majer, J. D. 1976. The maintenance of the ant mosaic in Ghana cocoa farms. Journal of Applied Ecology 13:123-144. McGlynn, T. P. 1999. The worldwide transfer of ants:
geographical distribution and ecological invasions.
Morton, S. R. and D. W. Davidson. 1988. Comparative structure of harvester ants communities in arid Austra- lia and North America. Ecological Monographs 58:19- 38.
Munger, J. C. 1992. Reproductive potential of colonies of desert harvester ants (Pogonomyrmex desertorum): effects of predation and food. Oecologia 90:276-282. Orr, M. R. 1992. Parasitic flies (Diptera: Phoridae) influence
foraging rhythms and caste division of labor in the leaf-cutter ant Atta cephalotes (Hymenoptera: Formicidae). Behavioral Ecology and Sociobiology 30:395-402.
Paine, R. 1968. A note on trophic complexity and community stability. American Naturalist 102:91-93.
Peeters, C. 1991. The ocurrence of sexual reproduction among ant workers. Biological Journal of the Linnean Society 44:141-152.
Porter, S. D. y M. A. Bowers. 1981. Emigration of an Atta colony. Biotropica 12:232.
Porter, S. D. y D. A. Savignano. 1990. Invasion of polygyne fire ants decimates native ants ad disrupt arthropod community. Ecology 71:2095-2106.
Porter, S. D., H. G. Fowler y W. P. MacKay. 1992. Fire ant mound densities in the United States and Brazil (Hymenoptera: Formicidae). Journal of Economic
Entomology 85:1154-1161.
Porter, S. D., H. G. Fowler, S. Campiolo y M. A. Pesquero. 1995a. Host specificity of several Pseudacteon (Diptera: Phoridae) parasites of fire ants (Hymenoptera: Formicidae) in South America. Florida Entomologist 78:70-75.
Porter, S. D., M. A. Pesquero, S. Campiolo y H. G. Fowler. 1995b. Growth and development of Pseudacteon phorid fly maggots (Diptera: Phoridae) in the heads of
Solenopsis fire ant workers (Hymenoptera: Formicidae). Environmental Entomology 24:475-479.
Porter, S. D., R. K. Vander Meer, M. A. Pesquero, S. Campiolo y H. G. Fowler. 1995c. Solenopsis (Hymenoptera: Formicidae) fire ant reactions to attacks of pseudacteon flies (Diptera: Phoridae) in southeastern Brazil. Annals
of the Entomological Society of America 88:570-575.
Rettenmeyer, C. W. 1962. The diversity of arthropods found with Neotropical army ants and observations on the behavior of representative species. Proceedings of the
North Central Branch of the Entomological Society of America 17:14-15.
Rettenmeyer, C. W., R. Chadab Crepet, M. G. Naumann y L. Morales. 1983. Comparative foraging by Neotropical army ants, pp.59-73 in: P. Jaisson, ed., Social Insects in
the Tropics Université Paris-Nord, Paris.
Rissing, S. W. 1987. Annual cycles in worker size of the seed- harvester ant Veromessor pergandei (Hymenoptera: Formicidae). Behavioral Ecology and Sociobiology 20:117-124.
Ryti, R. T. y T. J. Case. 1988a. Field experiments on desert ants: Testing for competition between colonies.
Ecology 69:1993-2003.
Ryti, R. T. y T. J. Case. 1988b. The regeneration niche of desert ants: Effects of established colonies. Oecologia 75:303-306.
Ryti, R. T. y T. J. Case. 1992. The role of neighborhood competition in the spacing and diversity of ant communities. American Naturalist 139:355-374. Savolainen, R. 1990. Colony succes of the submissive ant
Formica fusca within territories of the dominant Formica polyctena. Ecological Entomology 15:79-85.
Savolainen, R. y K. Vepsäläinen. 1988. A competition hierarchy among boreal ants: Impact on resource partitioning and community structure. Oikos 15:135- 155.
Schmidt-Hempel, P. 1992. Worker castes and adaptative demography. Journal of Evolutionary Biology 5:1-12. Schumacher, A. y W. G. Whitford. 1976. Spatial and temporal variation in Chihuahuan desert ant faunas.
Southwestern Naturalist 21:1-8.
Smallwood, J. 1982. Nest relocation in ants. Insectes Sociaux 29:138-147.
Stork, N. E. y T. M. Blackburn. 1993. Abundance, body size and biomass of arthropods in tropical forest. Oikos 67:483-489.
Sudd, J. H. y N. R. Franks. 1987. The Behavioral Ecology of
Ants. Blacwell, Glasgow.
Taber, S. W. 2000. Fire Ants. Texas University Press, Collegue Station.
Talbot, M. 1943. Population studies of the ant Prenolepis
imparis Say. Ecology 24:31-44.
Tennant, L. E. y S. D. Porter. 1991. Comparison of diets of two fire ants species (Hymenoptera: Formicidae): Solid and liquid components. Journal of Entomological Science 26:450-465.
Tobin, J. 1994. Ants as primary consumers: Diet and abundance in the Formicidae, pp. 297-307 in: J. H. Hunt y C. A. Napela, eds., Nourishment and Evolution in
Insect Societes Westview Press, Boulder.
Tobin, J. 1997. Competition and coexistence of ants in a small patch of rainforest canopy in peruvian Amazonia.
Journal of the New York Entomological Society
105:105-112.
Topoff, H. 1990. Slave making in ants. American Scientist 78:520-528.
Trivers, R. L. y H. Hare. 1976. Haplodiploidy and the evolution of the social insects. Science 191:249-263.
Tschinkel, W. R. 1991. Insect sociometry, a field in search of data. Insectes Sociaux 38:77-82.
Tschinkel, W. R. 1992. Brood raiding and the population dynamics of founding and incipient colonies of the fire ant, Solenopsis invicta. Ecological Entomology 17:179- 188.
Tschinkel, W. R. 1993. Sociometry and sociogenesis of colonies of the fire ant Solenopsis invicta during one annual cycle. Ecological Monographs 63:425-457. Tschinkel. W. R. y D. F. Howard. 1978. Queen replacement in
orphaned colonies of the fire ant, Solenopsis invicta.
Vander Meer, R. y L. Alonso. 1998. Pheromone directed behavior in ants, pp.159-192 in: R. Vander Meer, M. Breed, M. Winston y K. Espelie, eds., Pheromone
communication in Social Insects. Westview Press,
Boulder.
Ward, P. S. 1987. Distribution of the introduced argentine ant (Iridomyrmex humilis) in natural habitats of the Lowe Sacramento Valley and its effects on the indigenous ant fauna. Hilgardia 55:1-16.
Ward, P. S. 2000. Broad-scale patterns of diversity in leaf litter ant communities, pp.99-121 in D. Agosti, J. D. Majer, L. E. Alonso y T. R. Schultz, eds., Ants: Stan-
dard Methods for Measuring and Monitoring Biodiversity. Smithsonian Institution Press. Washing-
ton D.C.
Weber, N. A. 1972a. The Attines: The fungus culturing ants.
American Scientist 60:448-456.
Weber, N. A. 1972b. Gardening Ants: The Attines. American Philosophical Society, Philadelphia.
Wetterer, J. K. 1991. Allometry and the geometry of leaf- cutting ants in Atta cephalotes. Behavioral Ecology
and Sociobiology 29:347-351.
Whitcomb, W. H., A. Bhatkar y J. C. Nickerson. 1973. Predators of Solenopsis invicta queens prior to successful colony establishment. Environmental
Entomology 2:1101-1103.
Whitford, W. G. 1978. Structure and seasonal activity of Chihuahua desert ant communities. Insectes Sociaux 25:79-88.
Whitford, W. G. y G. Ettershank. 1975. Factors affecting foraging activity in Chihuahuan desert harvester ants.
Environmental Entomology 4:689-696.
Williams, D. F. 1994. Exotic ants: Biology, impact. And con-
trol of introduced species. Westview Press, Boulder.
Willis, E. O. 1983. A study of ant-following birds of northeastern. Brazil Reseach Reports of the National
Geographic Society 15:745-748.
Willis, E. O. y Y. Oniki. 1978. Birds and army ants. Annual
Review of Ecology and Systematics 9:243-263.
Wilson, E. O. 1958. Patchy distributions of ants species in New Guinea rain forests. Psyche 65:26-38.
Wilson, E. O. 1959. Some ecological characteristics of ants in New Guinea rain forests. Ecology 40:437-447. Wilson, E. O. 1961. The nature of the taxon cycle in the
Melanesian ant fauna. American Naturalist 95:169-193. Wilson, E. O. 1971. The Insect Societes Harvard University
Press, Cambridge.
Wilson, E. O. 1984. Tropical species parasites in the ant genus
Pheidole, with an analysis of the anatomical parasitic
syndrome (Hymenoptera: Formicidae). Insectes
Sociaux 31:316-334.
Wilson, E. O. 1985. The principle sof caste evolution.
Fortschritte der Zoologie 31:307-324.
Wilson, E. O. 1993. The Diversity of Life. W.W. Norton, New York.
Yanoviak, S. P. y M. Kaspari. 2000. Community structure and the habitat templet: ants in the tropical forest canopy and litter. Oikos 89:259-266.